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Vol.47, n. 3 : pp. 441-449, July 2004 ISSN 1516-8913 Printed in Brazil BRAZILIAN ARCHIVES OF BIOLOGY AND TECHNOLOGY AN INTERNATIONAL JOURNAL

Studies on the Morphometry of nobilii (, )

Pitchaimuthu Mariappan∗ and Chellam Balasundaram Aquaculture and Behaviour Unit - CRABU; Department of Science; Bharathidasan University; [email protected]; Tiruchirapalli; 620024; India

ABSTRACT

Fourteen morphometric characters of Macrobrachium nobilii (Henderson and Matthai, 1910), a freshwater prawn were measured from 72 males and 66 females collected at random in field. The growth constant (b) of total length and weight for male (4.28) and female (3.02) differed significantly (P<0.05). Comparison of slopes and elevations of regressions between sexes indicated significant difference in all the studied dependent parameters with carapace length (X), except with carapace width (Y). In males, major and minor chelipeds showed positive allometry while in females the growth was isometric.

Key words: Allometry, cheliped, freshwater prawn, length-weight, Macrobrachium nobilii

INTRODUCTION growth of chelipeds, especially that of Brachyuran has drawn detailed attention (Hartnoll, 1982), Allometric or relative growth pattern of various similar studies on individuals segments of decapod has been studied widely chelipeds are scanty (Jayachandran et al., 1996). (Hartnoll, 1978; Finney and Abele, 1981; The ratio between cheliped length and carapace Sumpton, 1990; Sarda and Cartes, 1997; Munio et length is species specific and this character has al., 1999), including genus Macrobrachium been subjected to detailed statistical analyses in (Holthuis, 1950; Koshy, 1973; Jayachandran, studies (Holthuis, 1950; Jayachandran, 1998; Mossolin and Bueno, 2003). Allometric 1998). In caridean prawns such as growth is determined by treating one variable (X) Macrobrachium, the chelipeds are taxonomically as reference or independent dimension and important (Chace and Bruce, 1993). Recently, comparing it with another dependent variable (Y) Suzuki and Kusamura (1997) differentiated M. (Snedecor and Cochran, 1967). In Crustaceans, nipponese from M. formosense on the basis of structures such as chelate legs in males, pleopod length and width of dactyl of second pereiopod. and abdominal width in females grow abruptly on Such investigations are also essential to establish attaining sexual maturity. This phenomenon is morphotypes in species such as M. rosenbergii. utilized to differentiate the mature and immature Moreover, the genus Macrobrachium contains population in many decapods, which are not more than 100 species; their identification is quite sexually dimorphic (Pinheiro and Fransozo, 1998; difficult due to great morphological variations Munio et al., 1999). Though the studies on the (Koshy, 1971). In India, the farmers still depend

∗ Author for correspondence

Brazilian Archives of Biology and Technology 442 Mariappan, P. and Balasundaram, C.

upon natural seed supply for stocking their ponds. minor chelipeds were individually weighed in an Hence, knowledge on the identity of the species electronic monopan balance (±0.001g). chosen for culture is an impelling necessity to Since sexual dimorphism is prominent in adult, M. eliminate mixing of species (Mariappan and nobilii, the data for males and females are Balasundaram, 1999). With this view, the study analyzed separately to fit regression lines to study was conducted to find out length-weight the allometeric growth pattern within and between relationship, growth pattern of chelipeds and sexes. In this study, the carapace length referred as segments of an endemic freshwater prawn CL was used as the reference dimension since as a Macrobrachium nobilii (Henderson and Matthai, taxonomic character CL was more reliable than 1910). Despite its small size, the culture of M. cephalothorax length (CTL) which often got nobilii is encouraged due to its commercial and damaged due to various factors (Grandjean et al., nutritional value (Mariappan et al., 2002; 2003) 1997). Crustacean allometric growth is defined by y=axb; this equation after log linearization yields log y = MATERIAL AND METHODS log a + b . x which eases the application of the results (Hartnoll, 1982). In the present study also A total of 72 males and 66 females of the data were log transformed before computing Macrobrachium nobilii were collected from Grand the regression equation (Sokal and Rohlf, 1981). Anicut, near Tiruchirpalli (10o 50’ N; 78o 43’ E) in In this context the derived slope value b referred as river Cauvery. It is the fourth largest river in allometric growth constant was designated as: 1) Indian Peninsula, which flows a distance of about isometric when 'b' ranged from 0.9 to 1.1 (Kuris 1171 kms. Total length, cephalo thoracic length, and Carlton, 1977), 2) positive if more than 1 and carapace width, carapace length, major and minor 3) negative if less than 1 (Tessier. 1960; Hartnoll, chela length, total weight of the animal and 1982). The variation between slopes and individual weight of chelae were calculated. elevations of the regression lines within and Length of the individual segments of major and between sexes were tested by analysis of minor chela was also measured (Fig. 1). covariance (Sndecor and Cochran, 1967). The variations among major and minor chela length and weight within and between sexes were tested separately.

RESULTS

Seventy-two males (total length: 46-77 mm) and 66 females (30-71 mm) were used for the morphometric study. Male M. nobilii had a minimum body weight of 1449 mg and a maximum of 12676 mg. The body weight of female ranged from 336 mg to 7178 mg. In

general, growth was manifested through size Figure 1 - Morphometry measurements of Macrobrachium nobilii. TL-Total length, CL-Carapace length, increment, which could be measured by increase CTL-Cephalothorax length, CW-Carapace in volume or weight (Kunju, 1978). Generally, the Width, MAC-Major Chela, MIC-Minor Chela. weight of crustaceans varies with the cube of the length (Jayachandran and Joseph, 1988). The increase in weight of both sexes of M. nobilii as The first two proximal segments i.e. coxa and measured by growth constant 'b' was higher for basis were ignored since their length was small males (4.28) than females (3.02), indicating the when compared to the total length of chelipeds positive allometric growth in this species. (see also Nagamine and Knight, 1980; Kuris et al., Subsequently, ANCOVA was performed to 1987). Measurements were taken to the nearest compare the slope and elevation of regression lines 0.1mm by using a Vernier caliber. Major and between males and females. The results indicated that there was a significant difference

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(P<0.001)(Table 1) between sexes for all the females (P>0.05)(Table 2). For both sexes the ratio chosen variables except for carapace width and between major and minor chela length and weight total weight (P>0.05). In males, the body weight, were calculated. The length ratio varied from 1.15 chelae length and weight with reference to to 1.58 and that of weight varied from 0.87 to 2.42 carapace length indicated positive allometric for males. For females, the chela length ranged growth while other parameters such as total length, from 1 to 1.43 and the weight ratio ranged from 1 cephalothorax length and carapace width showed to 2.08. The change in the chela length (major negative allometry (Table 1). /minor) remained constant (P>0.05) in male and female. A similar trend was observed with Chela Growth reference to major and minor chela weight ratio In females, the growth pattern of major and minor for females. However, for males there was a chela length was isometric. In males, there was a significant inverse relationship between major remarkable difference in the pattern of length chela weight/minor chela weight ratio and increment between major and minor chela carapace length. (P<0.001) but no such difference prevailed either in chela weight in males or length and weight of

Table 1 - Macrobrachium nobilii: Analysis of Covariance (ANCOVA) to test the slopes (b) and elevations of regression equation for males (n=72) females (n=66). Characters Regression Equation Comparison of Allometry Male Female Slope Elevation Male Female Growth in relation to carapace length Total length 1.022+0.608.X 0.87+0.702.X 1.61* 237.53*** - - Cephalothorax length 0.359+0.862.X 0.291+0.885.X 10.44*** 40.59*** - - Carapace width -0.011+0.862.X -0.054+0.886.X 0.7 NS 75.25*** - - Major chela length -0.534+1.92.X 0.395+0.972.X 32.28*** 109.15*** + 0 Minor chela length -0.61+1.88.X 0.424+0.905.X 37.12*** 157.06*** + 0 Major chela weight -3.074+4.51.X -0.994+2.388.X 25.18*** 236.18*** + + Minor chela weight -4.255+5.371.X -1.176+2.44.X 49.17*** 293.86*** + + Chela weight -3.264+4.865.X -0.76+2.4.X 53.35*** 323.03*** + + Total weight 8.022+2.929.X 5.012+2.853.X 0.10 NS 193.87*** + + * Statistically significant (P<0.05): F70,64 = 1.502 *** Statistically significant (P<0.001): F70,64 = 2.166 NS Statistically not significant (P>0.05): F70,64

Table 2 - Macrobrachium nobilii: Summary of Analysis of Covariance results obtained between major and minor chelae length and weight in relation to carapace length. Variations Reduction Df SS MS F Male chela length Between chelae 1.95 70 4.21 0.06 8.36*** Within chelae 1.88 70 0.50 0.007 Male chela weight Between chelae 10.82 70 2.05 0.029 0.75NS Within chelae 15.34 70 2.74 0.039 Female chela length Between chelae 0.55 64 0.46 0.007 1.49NS Within chelae 0.48 64 0.31 0.005 Female chela weight Between chelae 3.32 64 3.18 0.049 1.28NS Within chelae 3.48 64 2.49 0.039 *** Statistically Significant (P<0.001): F70,70 = 2.116 NS Statistically not significant (P>0.05): F70.70 = 1.485 for males F64,64 = 1.513 for females.

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Table 3 - Regression equations for individual segments of major and minor chela in relation to carapace length in male and female. Parameter R Intercept (a) Slope (b) Allometry Male-Major Chela Ischium 0.878*** -0.784 1.366 + Merus 0.855*** -0.133 1.833 + Carpus 0.829*** -1.344 2.145 + Propodus 0.839*** -1.174 2.104 + Dactylus 0.852*** -1.129 1.632 + Male-Minor Chela Ischium 0.789*** -0.532 1.116 + Merus 0.764*** -0.708 1.412 + Carpus 0.767*** -0.857 1.627 + Propodus 0.706*** -0.636 1.567 + Dactylus 0.703*** -0.493 1.049 + Female-Major Chela Ischium 0.818*** -0.061 0.681 - Merus 0.867*** -0.256 0.961 0 Carpus 0.832*** -0.370 1.145 + Propodus 0.812*** -0.113 1.04 + Dactylus 0.872*** -0.336 0.859 - Female-Minor Chela Ischium 0.803*** -0.014 0.614 - Merus 0.851*** -0.27 0.927 0 Carpus 0.883*** -0.272 0.998 0 Propodus 0.863*** -0.089 0.957 0 Dactylus 0.821*** -0.256 0.746 - + Positive allometry, - Negative allometry, 0 isometry *** Highly significant (P<0.001)

Segmental growth of major and minor chela with (P>0.05). However elevations (intercept of minor reference to the carapace length of M. nobilii was and major chela) differed significantly (P<0.001) analyzed among sexes. Among the five segments, (Table 4). Regarding the growth of segments of propodus was the longest segment followed by major and minor chelae in male and female, the carpus and merus in both chelae of male and in length increment of each segment differed one female; ischium and dactylus was the smallest from another (P<0.001) (Table 5). segments. In male all the segments in major and Among sexes, carpus had a maximum allometry minor chela showed positive allometric growth level, which declined to the minimum in dactylus (Table 2). and ischium (Fig. 2). In female, carpus and propodus of major chela only showed positive isometric growth. Further merus of major and minor chela exhibited isometric growth. Growth of carpus and propodus of minor chela was also isometric. Ischium and dactylus in major and minor chelae showed negative allometric growth (Table 3). Analysis of covariance of growth of major and minor chelae segments indicated that there was a significant difference (P<0.001) in growth pattern between major and minor chelae in males whereas in females the growth constants of major and minor chela segments did not differ significantly

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2.5 MMC MMiC FMC FMiC

2

1.5

1 Allometry Level 0.5

0 Ischium Merus Carpus Propodus Dactylus Chela Segment

Figure 2 - Allometry levels of major and minor chela segments of male and female Macrobrachium nobilii. MMC-Male Major Chela, MMiC-Male Minor Chela, FMC-Female Major Chela, FMiC-Female Minor Chela.

Table 4 - Macrobrachium nobilii: Comparison or regression equations for individual segments (Y) of major and minor chela against carapace length (X) in males. Character Regression Equation Comparison of Major Minor Slope Value Elevation Value Male Ischium -0.783+1.366.X -0.532+1.116.X 3.37*** 164.89*** Merus -1.133+1.833.X -0.708+1.412.X 5.32*** 171.31*** Carpus -1.344+2.146.X -0.857+1.627.X 4.59*** 198.47*** Propodus -1.174+2.104.X -0.636+1.567.X 5.23*** 127.38*** Dactylus -1.129+1.632.X -0.493+1.049.X 6.73*** 38.69*** Female Ischium -0.611+0.681.X -0.014+0.613.X 0.29 NS 61.84** Merus -0.256+0.961.X -0.27+0927.X 0.11 NS 156.94** Carpus -0.369+1.144.X -0.272+0.998.X 1.61* 164.56** Propodus -0.113+1.039.X -0.089+0.957.X 0.5 NS 164.91** Dactylus -0.335+0.858.X -0.256+0.747.X 1.59* 160** * Statistically significant (P<0.05): F64,64 = 1.513 ** Statistically highly significant (P<0.001): F64,64 = 2.193 *** Statistically highly significant (P<0.001): F70,70 = 2.116 NS Statistically not significant (P>0.05): F64,64 = 1.513

Brazilian Archives of Biology and Technology 446 Mariappan, P. and Balasundaram, C.

Table 5 - Macrobrachium nobilii: Comparison of regression equations for individual segments (Y) of major of male against female and minor chela of male against female carapace length (X). Character Regression Equation Comparison of Male Female Slope Value Elevation Value Major Chela Ischium -0.783+1.366.X -0.611+0.681.X 42.15*** 196.6*** Merus -1.133+1.833.X -0.256+0.961.X 36.44*** 241.31*** Carpus -1.344+2.146.X -0.369+1.144.X 27.27*** 250.09*** Propodus -1.174+2.104.X -0.113+1.039.X 53.86*** 275.56*** Dactylus -1.129+1.632.X -0.335+0.858.X 19.63*** 78.31*** Minor Chela Ischium -0.532+1.116.X -0.014+0.613.X 9.36*** 98.66*** Merus -0.708+1.412.X -0.27+0927.X 10.22*** 244.32*** Carpus -0.857+1.627.X -0.272+0.998.X 14.86*** 253.91*** Propodus -0.636+1.567.X -0.089+0.957.X 10.87*** 253.79*** Dactylus -0.493+1.049.X -0.256+0.747.X 4.87*** 247.18*** *** Statistically highly significant (P<0.001) F70, 64 = 2.166

DISCUSSION females during courtship in male Corystes (Hartnoll, 1968), inter male combat in non Length-weight relationship study is a useful index brachyurans like Caprella gorgonia (Lewbel, to measure the variation in the growth of 1978); Macrobrachium rosenbergii (Barki, 1991), individual prawn or group of prawns M. australiense (Lee and Fielder, 1983), burrow (Jayachandran and Joseph, 1988). Variation in the protection in Alpheus heterochaelis (Conover and length weight relationship obtained between the Miller, 1978); holding the female during male and female of M. nobilii was due to the copulation in Orconectes propinquus (Stein, 1976) variation in food, population density, and protecting the defenseless female just environmental conditions and genetic make up of after premating moult in M. nobilii the species which has been well documented in (Balasundaram, 1980). earlier studies (Abrahamson, 1966; Kuris et al., Among the closely related species of 1987). In decapod crustaceans, morphometric Macrobrachium such as M. malcolmsonii and M. studies have been widely used for taxonomic birmanicum, though the ratio between merus and purposes in the genus Macrobrachium (Tesch, palm, carpus and chela, and fingers and palm were 1915; Holthuis, 1950; Koshy, 1969; Jayachandran the same, in M. malcolmsonii, the carpus was and Joseph, 1988; Chace and Bruce, 1993; Suzuki longer than the palm while it is shorter in M. and Kusamura, 1997); in brachyura (Davidson and birmanicum (Jayachandran, 1998). In M. nobilii, Marsden, 1987; Clayton, 1990; Abello et al., the carpus was shorter than the propodus while the 1990); in spiny and clawed lobsters (Aiken and level of allometry was higher when compared to Waddy, 1980). It is well known that propodus. Koshy (1973) and Nagamine and Macrobrachium is a more confusing genus. Knight (1980) also made similar findings in M. Recently, Jayachandran (1998) differentiated some dayanum and M. rosenbergii, respectively where closely related species of Macrobrachium on the the propodus was longer than the carpus. In M. basis of the relative growth of second pereiopods. rosenbergii, three male morphotypes (i.e) BC-Blue Like other crustacean groups the growth of male Clawed Males, OC-Orange Clawed Males and cheliped had a positive allometry with reference to SM-Small Males were differentiated on the basis the cephalothoracic length but in females it was of the total length of chela (Kuris et al., 1987). isometric. Such variations in allometric level are Further BC males were distinguished from OC well documented in Brachyura. The enlarged males from the variations in length of carpus and chelae in males are related to the vital social propodus. The propodus had a positive allometric interactions like territorial defense, combat, (β=1.26) while the growth of carpus was isometric display and courtship. The use of such enormous (β=1.09)(Kuris et al., 1987). In male M. nobilii, sized chelae in decapods are many viz; signaling the growth of propodus and carpus had positive display by male Uca (Crane, 1975), carrying allometry. In females it was isometry for major

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and minor chela. Comparison of allometric levels RESUMO in different crustacean species were indicated, its variations were as a function of sex and even Quatorze características morfológicas do within females of Trapezia ferruginea, allometric Macrobrachium nobilii (Henderson e Matthai, level was negative in normal while it was positive 1910), 72 machos e 66 fêmeas de lagostas de água among berried females (Finney and Abele, 1981). fresca foram medidas. O crescimento constante (b) Huxely (1932) developed a growth gradient do comprimento total e peso para os machos (4.28) concept by studying the growth pattern of a series e fêmeas (3.02) apresentaram diferenças of organs or segments of an organ. In this line the significativas (P<0.05). A comparação da best known examples are the studies on the inclinação e da elevação das regressões entre sexos patterns of growth of a) individual segments of a indicaram diferenças significativas em todos os limb, b) the segments of the body and c) the parâmetros dependentes estudados comprimento sequence of limbs along the body (Hartnoll, 1982). da carcaça (x), exceto largura da carcaça (y). Nos Huxley (1932) observed a growth center in machos, as garras principais e secundárias chelipeds of species like Uca pugnax, Maja demonstraram alometria positiva quando nas squidao and Macrobrachium rude (=Palaemon fêmeas o crescimento foi isométrico. rudis) where the maximum allometry level was at propodus, which decreased towards dactylus and coxa. Dawes (1934) observed such a growth center REFERENCES in propodus of Alpheus dentipes but Asellus aquaticus it was in propodus or carpus (Needham, Abello, P.; Pertierra, J. P. and Reid, D. G. (1990), 1943). In Ocypode aegyptica and Eriocheir Sexual dimorphism, relative growth and handedness sinensis two growth centers have been reported in Liocarcinus depurator and Macropipus (Sandon, 1937). In the right chela of male Pagurus tubercilatus (Brachyura: Portunidae). Scientia bernhardus the growth gradient changed from the Marina, 54,195-202. merus to propodus when the crab grows (Huxley, Abrahamsson, S. (1966), Dyanamics of an isolated 1932). In this study the maximum allometry level population of the crayfish Astacus astacus (Linne) and Pasifastacus leniusculus (Dana). Svensk in M. nobilii was recorded in carpus in both chelae Naturvetenskap, 1964, 306-316. and both sexes. Presence of sexual dimorphism in Aiken, D. E. and Waddy, S. L. (1980), Maturity and second pereiopod in M. nobilii was an agreement Reproduction in the American lobster. Can. Tech. with the earlier findings in related species such M. Rep. Fish. Aquat. Sci., 932, 59-71. rosenbergii (Nagamine and Knight, 1980), M. Balasundaram, C. (1980), Ecophysiological studies in malcolmsonii (=Palaemon malcalmsonii) prawn culture (Macrobrachium nobilii). Ph.D. (Patwardhan, 1937) and M. dayanum (Koshy, Dissertation - Madurai Kamaraj University, India. 1973). From this study we concluded that the Barki, A.; Karplus, I. and Goren, M. (1991), studied morphometric characters could be used to Morphotype related dominance hierarchies in males describe the species. of Macrobrachium rosenbergii (Crustacea, Palemonidae). Behaviour, 117, 145-160. Chace, F. A. and Bruce, A. J. (1993), The caridean shrimps (Crustacea, Decapoda) of the Albatross ACKNOWLEDGEMENTS Philippine Expedition, 1907-1910, part 6 Superfamily Palaemonidae. Smithsonian Contribution to Zoology, PM is grateful to CSIR-India for the award of 543, 1-152. Senior Research Fellowship and Senior Research Clayton, D. A. (1990), Crustacean allometric growth: a Fellowship (Extended), Prof. K.V. Jayachandran, case for caution. Crustaceana, 58, 270-290. Kerala Agricultural University for his critical Conover, M. R. and Miller, D. E. (1978), The review of this MS and P. Balamurugan for his importance of large chela in the territorial and pairing behaviour of the snapping shrimp, Alpheus valuable helps in MS preparation. heterochaelis. Mar. Behav. Physiol., 5, 185-192. Crane, J. (1975), The fiddler crabs of the world (Ocypodidae: Genus Uca). New Jersey : Princeton University Press.

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Received: May 29, 2002; Revised: February 18, 2003; Accepted: October 16, 2003.

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